Levels of behavioral organization and the evolution of division of labor

The major features of insect societies that fascinate biologists are the self-sacrificing altruism expressed by colony members, the complex division of labor, and the tremendous plasticity demonstrated in the face of changing environments. The social behavior of insects is a result of complex interactions at different levels of biological organization. Genes give rise to proteins and peptides that build the nervous and muscular systems, regulate their own synthesis, interact with each other, and affect the behavior of individuals. Social behavior emerges from the complex interactions of individuals that are themselves far removed from the direct effects of the genes. In order to understand how social organization evolves, we must understand the mechanisms that link the different levels of organization. In this review, we discuss how behavior is influenced by genes and the neural system and how social behavior emerges from the behavioral activities of individuals. We show how different levels of organization share common features and are linked through common mechanisms. We focus on the behavior of the honey bee, the best studied of all social insects.